Journal of Microbiology (2014) Vol. 52, No. 2, pp. 179–183
1007/s12275-014-2720-5
Copyright G2014, The Microbiological Society of Korea
NOTE
Effects of Light Intensity on Components and Topographical Structures
of Extracellular Polysaccharides from the Cyanobacteria Nostoc sp.
Hongmei Ge1,2, Ling Xia1,2, Xuping Zhou1,2,
3 1
Delu Zhang , and Chunxiang Hu *
1
Key Laboratory of Algal Biology, Institute of Hydrobiology,
Chinese Academy of Sciences, Wuhan, 430072, P. R. China
2
Graduate University of Chinese Academy of Sciences, Beijing, 100049,
P. R. China
3
Department of Biological Science and Biotechnology, Wuhan University
of Technology, Wuhan, 430070, P. R. China
(Received Dec 31, 2012 / Revised May 8, 2013 / Accepted Jul 30, 2013)
A study on the effects of light intensity (40 and 80 μE/m2/sec)
on the components and topographical structures of extra-
cellular polysaccharides (EPS) was carried out in cyanobac-
teria Nostoc sp.. EPS yield increased with light intensity.
However, light intensity did not significantly affect the EPS
fractions and monosaccharide composition. Higher light
intensity generally resulted in higher protein content of EPS
in similar fractions. The topographical structure of EPS, in-
vestigated by atomic force microscopy, appeared as spherical
lumps, chains and networks. The long chains were observed
at higher light intensity. Thus, light intensity affected the
yield and nature of EPS.
Keywords: cyanobacteria, extracellular polysaccharides, light
intensity, Nostoc sp., atomic force microscopy
Many cyanobacterial species can synthesize and secrete ex-
tracellular polysaccharides (EPS), which contained amino
acids, polypeptides, amides, proteins, vitamins, especially
polysaccharides (Neu and Marshall, 1990). Cyanobacterial
EPS can be divided in two main groups: the polysaccharides
released into the surrounding environment (RPS), and the
capsular polysaccharides associated with the cell surface
(CPS), which can be referred to as sheaths, capsules, and
slimes (De Philippis and Vincenzini, 1998, 2003). EPS is
increasingly gaining particular research attention because
of its potentially useful applications (De Philippis and Vin-
cenzini, 1998, 2003; De Philippis et al., 2001, 2011; Suresh
Kumar et al., 2007).
Various culture parameters affect the synthesis of EPS
(De Philippis and Vincenzini, 1998; Pereira et al., 2009). In
particular, light is one of the most important factors that
*For correspondence. E-mail: cxhu@ihb.ac.cn; Tel.: +86-27-68780866
DOI 10.
influence the synthesis and release of EPS. Several recent
studies on the influence of light intensity (Friedman et al.,
1991; Moreno et al., 1998; Otero and Vincenzini, 2003;
Trabelsi et al., 2009; Yu et al., 2010; Mota et al., 2013) and
wavelengths (Ehling-Schulz et al., 1997; Chen et al., 2009)
on EPS yield have been reported. Different light regimens
(continuous light and light-dark cycles) do not significantly
affect the monosaccharidic composition of EPS (Vincenzini
et al., 1993; De Philippis and Vincenzini, 1998). However,
little information is available about the effects of light in-
tensity on the monosaccharidic composition of EPS.
Nostoc sp. is a exopolysaccharide-secreting cyanobacterium
dominant in biological soil crusts (BSCs) (Hu et al., 2003).
This study aims to investigate the effect of light intensity
on EPS component from Nostoc sp. and the effect of light
intensity on morphological structure of the EPS using atomic
force microscopy (AFM). A physicochemical investigation
would provide essential information on the function and
potential uses of this polysaccharides.
Nostoc sp. (FACHB 892) was separated from BSCs from
Tengger Desert, China and conserved in the Institute of
Hydrobiology, Chinese Academy of Sciences. Cultures were
carried out axenically in 250 ml glass flasks containing 200 ml
of BG-11 medium (Rippka et al., 1979) at 25±1°C and were
continuously illuminated laterally on one side by a combi-
nation of cool-white. Light intensity was controlled at 40 and
80 μE/m2/s with a quantitherm light meter thermomter
(Hansatech, UK). Cultures were inoculated at a chlorophyll
a concentration of 1.4 μg/ml.
Chlorophyll a fluorescence was measured using a plant ef-
ficiency analyzer (Hansatech). Nostoc sp. was dark-adapted
for 15 min before measuring the fluorescence parameter
Fv/Fm (Photosystem II activity). The saturating light pulse
was 1,500 μE/m2/s.
After 20 days of growth, the cells in suspension were har-
vested by filtration and freeze-dried. Biomass measured as
dry cell weight. The polysaccharides content released into the
culture medium (RPS), in the supernatant, was determined
according to the procedure of Chen et al. (2003). The poly-
saccharides formed capsular or slimy layer (CPS) were pre-
pared using the method of Su et al. (2008). The amount of
total exopolysaccharides (EPS) were the sum of the amounts
of RPS and CPS.
Purification of RPS was performed according to the pro-
cedure of Hu et al. (2003). The filtered supernatants were
concentrated in a rotary evaporator (BUCHI Rotavapor
R-210, Switzerland) at 35°C. The concentrates were directly
180 Ge et al.
Fig. 1. Effect of different light intensities on Fv/Fm of Nostoc sp.
placed in a DEAE-Sepharose fast-flow column (5×50 cm).
The column was first eluted with distilled water for collec-
tion of the neutral polymers, then eluted with 1.0 M NaCl
to collect the acidic I polymers, followed by 2.0 M NaCl to get
the acidic II polymers. For CPS, the supernatants extracted
from dry cells maintained at 80°C for 6 h were prepared us-
ing the same method as that of RPS.
Carbohydrate and protein contents of the purified EPS
were measured using the phenol-sulfuric acid method
(Dubois et al., 1956) and Lowry’s method (Lowry et al.,
1951), respectively.
Quantitative determination of the carbohydrate composi-
tion was carried out according to Hokputsa et al. (2003).
The trimethysilylated samples were analyzed by gas chro-
matography (Agilent 6890, USA) with a DB-5 fused silica
capillary column. The temperature program was as follows:
the initial column temperature of 150°C was held for 1 min;
the temperature was then increased to 220°C at a rate of
4°C/min; held at 220°C for 10 min, and then increased
again to 250°C at a rate of 10°C/min and held for 5 min.
The temperature of the injector was 250°C. The split ratio
was 30:1; nitrogen was used as the carrier gas, and the flow
rate of N2 was 0.5 ml/min. The temperature of the flame
ionization detector was 280°C.
Samples for AFM were prepared as follows: the purified
EPS fraction was dissolved in ultrapure water (1 mg/ml),
Table 1. The amount of EPS from Nostoc sp. and the percentage of one neutral and two acidic fractions of purified EPS (% of total purified EPS content)
under different light intensities
Biomass (g DW/L)
Yield (mg/g DW) RPS
CPS
EPS
Percentage (%) RPS Water eluates
1.0 M NaCl eluates
2.0 M NaCl eluates
CPS Water eluates
1.0 M NaCl eluates
2.0 M NaCl eluates
DW indicates dry weight of biomass.
stirred and centrifuged for 30 min at 3,000×g; the super-
natant was diluted to a final concentration of 10-3 mg/ml. The
diluted EPS solutions were spread on freshly cleaved mica
disks and fixed for 2 min, and then dried with N2 gas. AFM
(Agilent 5500, USA) was carried out in air at 25±0.5°C and
50–60% relative humidity. All imagines were obtained in
tapping mode. Three samples were text for each figure. The
molecular height and length on AFM images were meas-
ured using the AFM analysis software.
Analysis of variance was performed using the SPSS 18.0,
with the significance level set at 0.05 or 0.01. Values are ex-
pressed as the mean±standard deviation (n=3).
After 20 days of culture, the Fv/Fm of Nostoc sp. at 40
μE/m2/s was 1.1-fold higher than that at 80 μE/m2/s (P<0.05)
(Fig. 1). The biomass of Nostoc sp. at 40 and 80 μE/m2/s
did not differ significantly (P>0.05) (Table 1). These results
suggest that 40 μE/m2/s was more optimal for Nostoc sp.
culture compared with 80 μE/m2/s.
The yields of RPS, CPS and EPS produced by Nostoc sp. at
80 μE/m2/s (134.26, 71.94 and 206.20 mg/g DW for RPS,
CPS and EPS, respectively) were all higher than that at 40
2
μE/m /s (97.78, 57.70, and 155.49 mg/g DW) (P<0.01)
(Table 1). Our data are in agreement with previous studies
that EPS production is enhanced by high light intensities
(Friedman et al., 1991; Moreno et al., 1998; Otero and
Vincenzini, 2003; Trabelsi et al., 2009; Yu et al., 2010; Mota
et al., 2013), suggesting that energy availability significantly
affects the EPS biosynthesis of cyanobacteria. Higher light
intensity results in higher EPS production, which may con-
stitute mechanism of releasing the excess energy absorbed
by the cells (Mota et al., 2013) and increasing CO2 fixation
rate in the cells under high light intensity.
The polysaccharides produced by Nostoc sp. were separated
into three parts, one neutral and two acidic fractions by
anion exchange chromatography. The percentages of each
fraction are listed in Table 1. The 2.0 M NaCl eluates
(7.36–9.50%) only accounted for a small proportion of the
total; thus, were not characterized further. Light intensity
did not significantly affect the fractions of RPS or CPS
from Nostoc sp. The water eluates comprised the main
fraction of RPS, whereas the 1.0 M NaCl eluates comprised
the main fraction of CPS (Table 1). Results from a previous
study revealed that the water eluates comprised the main
fraction of CPS in three soil Nostoc species (Huang et al.,
2 2
40 μE/m /s 80 μE/m /s
0.82±0.00 0.83±0.05
97.78±1.38 134.26±7.03
57.70±5.08 71.94±3.95
155.49±6.46 206.20±10.98
56.19±1.23 57.97±2.50
34.31±1.68 34.67±2.27
9.50±0.44 7.36±0.24
16.74±0.39 15.30±1.00
74.90±1.04 76.40±1.40
8.37±0.65 8.30±0.39
 Effects of light intensity on EPS components and topographical structures 181

Table 2. Monosaccharide composition (% of total carbohydrate content) and the total carbohydrate content and total protein content (% of poly-
saccharides dry weight) of the purified EPS of Nostoc sp. under different light intensities
40-W 40-1 80-W 80-1
RPS Rhamnose 6.5±0.2 3.5±0.0 9.6±0.2 8.7±0.2
Xylose 9.1±0.4 6.4±0.1 6.3±0.1 4.6±0.1
Mannose 4.5±0.1 2.2±0.0 8.3±0.1 3.6±0.0
Galactose 10.5±0.2 12.4±0.1 13.1±0.0 16.1±0.1
Glucose 63.2±0.9 71.5±0.0 54.2±0.3 59.4±0.2
Galacturonic acid 3.7±0.1 3.9±0.0 5.2±0.1 7.7±0.2
Glucuronic acid 2.4±0.0 - 3.3±0.0 -
Total carbohydrate 58.6±1.3 52.8±2.2 62.2±4.5 38.0±2.2
Total protein 11.3±1.2 11.5±0.9 16.1±0.5 19.2±1.7

CPS Rhamnose 1.5±0.0 13.8±0.1 - 10.8±0.1

Xylose - 5.4±0.0 1.2±0.0 6.0±0.1
Mannose 2.2±0.0 6.9±0.1 1.9±0.0 7.3±0.2
Galactose 23.4±0.1 12.5±0.3 22.2±0.0 13.8±0.5
Glucose 62.9±0.2 49.8±0.4 63.9±0.0 53.5±0.7
Galacturonic acid 4.0±0.0 3.3±0.0 4.4±0.0 2.9±0.1
Glucuronic acid 6.3±0.7 8.4±0.5 6.3±0.0 5.7±1.5
Total carbohydrate 56.4±2.3 29.8±2.1 66.6±5.3 29.6±2.1
Total protein 18.7±1.4 38.0±1.7 15.3±1.1 48.3±1.9
40-W, water eluates from 40 μE/m2/s; 40-1, 1.0 M NaCl eluates from 40 μE/m2/s; 80-W, water eluates from 80 μE/m2/s; 80-1, 1.0 M NaCl eluates from 80 μE/m2/s.

1998). Hu et al. (2003) reported that the 1.0 M NaCl elu-
ates were the main fraction of RPS from a soil algae Nostoc
sp. In the present study, the difference in the main fractions
between RPS and CPS suggests that the physiological proper-
ties between RPS and CPS were different.
The contents of purified EPS carbohydrates and proteins
are shown in Table 2. For RPS, light did not significantly
affect the carbohydrate content in the water eluates (P>0.05);
whereas the content of carbohydrate in 1.0 M NaCl eluates
at 40 μE/m2/s was 1.4-fold higher than that at 80 μE/m2/s
2
(P<0.01). And the RPS protein contents at 80 μE/m /s
(16.1% of polysaccharides and 19.2% of polysaccharides for
water eluates and 1.0 M NaCl eluates, respectively) were
2
higher than that at 40 μE/m /s (11.3%, 11.5%) (P<0.01)
(Table 2). For CPS, the carbohydrate content of water
elutes at 80 μE/m2/s was 1.2-fold higher than that at 40
2
μE/m /s (P<0.05); whereas the carbohydrate contents were
similar for the said light intensities in 1.0 M NaCl eluates
(P>0.05). And the protein content of 1.0 M NaCl eluates at
2 2
80 μE/m /s was 1.3-fold higher than that at 40 μE/m /s (P
<0.01) (Table 2). Thus, the EPS protein content at 80 μE/m2/s
was generally higher than that at 40 μE/m2/s in similar
fractions (P<0.01).
The principal components of EPS are polysaccharides or
proteoglycans, wherein the protein part is covalently linked
to carbohydrate (Hu et al., 2002). In this study, the rate of
nitrate assimilation and metabolism in Nostoc sp. may be
higher at higher light intensity. Furthermore, proteins can
be glycosylated to form glycoproteins, which can be trans-
located to the cell surface or into the surrounding media

(A) (B) (C) (D) Fig. 2. AFM images of purified EPS

from Nostoc sp. under different light
intensities (image size = 2×2 μm).
For RPS: (A) 40-W, (B) 40-1, (C)
80-W, (D) 80-1. For CPS: (E) 40-W,
(F) 40-1, (G) 80-W, (H) 80-1. 40-
2
W: water eluates from 40 μE/m /s;
40-1: 1.0 M NaCl eluates from 40
2
μE/m /s; 80-W: water eluates from
2
80 μE/m /s; 80-1: 1.0 M NaCl elu-
ates from 80 μE/m2/s.

(E) (F) (G) (H)

182 Ge et al.
Table 3. Molecular sizes of EPS molecules by AFM
Spherical lump
Height range (nm)
RPS 40-W 8.11–15.88
40-1 6.08–11.15
80-W 4.95–16.17
80-1 6.42–16.56
CPS 40-W 5.07–9.8
40-1 9.46–28.05
80-W 4.05–17.91
80-1 - -
2 2
40-W, water eluates from 40 μE/m /s, 40-1: 1.0 M NaCl eluates from 40 μE/m /s; 80-W, water eluates from 80 μE/m /s; 80-1, 1.0 M NaCl eluates from 80 μE/m /s.
(Pereira et al., 2009). Thus, the protein content of EPS was
higher at high light intensity.
Light intensity did not significantly affect the monosac-
charide composition of RPS or CPS (Table 2). In the carbo-
hydrate part of RPS or CPS, glucose and galactose were the
most abundant, rhamnose, xylose, mannose, galacturonic
acid, and glucuronic acid were present as minor components.
Different light regimens did not affect the monosaccharidic
composition of EPS (Vincenzini et al., 1993; De Philippis
and Vincenzini, 1998). We found that light intensity also
did not significantly affect the quality of the polymer des-
cribed above. For most algal strains, RPS and CPS contain
the same sugar composition, suggesting that RPS could ori-
ginate from CPS (Li et al., 2001; Bellezza et al., 2003; Pereira
et al., 2009; Yoshimura et al., 2012). Our results are in agree-
ment with previous studies.
The topographical AFM images of EPS are presented in
Fig. 2, and the molecular sizes of EPS are shown in Table 3.
For RPS, the height of short chain in water eluates at 40
μE/m2/s (10.14–17.23 nm) was higher than that at 80 μE/m2/s
(4.29–8.58 nm) (P<0.01) (Fig. 2A, 2C, and Table 3). And in
1.0 M NaCl eluates, the short chains were observed at 80
μE/m2/s, compared with that at 40 μE/m2/s (Fig. 2B, 2D
and Table 3). For CPS, the long chains were observed at 80
2 2
μE/m /s, compared with the corresponding 40 μE/m /s re-
sults in similar fractions (Fig. 2E, 2F, 2G, 2H, and Table 3).
Clearly, the 1.0 M NaCl eluates at 80 μE/m2/s appeared as
networks, which were thought to contain long chains (Fig.
2H).
Different EPS molecules have different topographical struc-
tures. Pletikapic et al. (2011) confirmed the surface mor-
phology of EPS isolated from Cylindrotheca closterium ap-
peared as networks at 10 μg/ml. The AFM image of EPS
from Amphidinium carterae appeared as different particle
sizes, with 181 nm average roughness (Mandal et al., 2011).
The present study on the EPS from Nostoc sp. under diffe-
rent light intensities showed spherical lumps, chains and
networks. From the AFM images (Fig. 2), the topographical
structures of the exopolysaccharides in similar fractions
under different light intensities were evidently different, sug-
gesting that light intensity affected the physicochemical
properties of EPS from Nostoc sp.
In conclusion, the results obtained in this work show that
higher light intensity lead to higher EPS yield produced by
Nostoc sp. The analysis of the purified EPS revealed seven
types of monosaccharide, in which glucose and galactose
Chain Net work
Length range (nm) Height range (nm) Height range (nm)
280–630 10.14–17.23 -
- - -
420–520 4.29–8.58 -
400–400 14.53–18.25 -
- - -
- - -
780–1040 6.08–17.23 -
- 6.76–15.54
2 2
were the most dominant. Moreover, the contents of EPS
carbohydrate and protein in similar fractions were different,
suggesting light intensity affects the EPS components. The
AFM results further indicate that light intensity affects the
physicochemical properties of EPS from Nostoc sp.
This study was supported by the National Natural Science
Foundation of China (Nos. 31170464; 30870470) and the
National Forestry Public Welfare Industry Research Project
(Nos. 201404204).
References
Bellezza, S., Paradossi, G., De Philippis, R., and Albertano, P. 2003.
Leptolyngbya strains from Roman hypogea: cytochemical and
physico-chemical characterisation of exopolysaccharides. J. Appl.
Phycol. 15, 193–200.
Chen, L., Li, D., and Liu, Y. 2003. Salt tolerance of Microcoleus vagi-
natus Gom., a cyanobacterium isolated from desert algal crust,
was enhanced by exogenous carbohydrates. J. Arid Environ. 55,
645–656.
Chen, L., Wang, G., Hong, S., Liu, A., Li, C., and Liu, Y. 2009. UV-
B-induced oxidative damage and protective role of exopolysac-
charides in desert cyanobacterium Microcoleus vaginatus. J.
Integr. Plant Biol. 51, 194–200.
De Philippis, R., Colica, G., and Micheletti, E. 2011. Exopolysac-
charide-producing cyanobacteria in heavy metal removal from
water: molecular basis and practical applicability of the bio-
sorption process. Appl. Microbiol. Biotechnol. 92, 697–708.
De Philippis, R., Sili, C., Paperi, R., and Vincenzini, M. 2001.
Exopolysaccharides-producing cyanobacteria and their possible
exploitation: a review. J. Appl. Phycol. 13, 293–299.
De Philippis, R. and Vincenzini, M. 1998. Exocellular polysaccha-
rides from cyanobacteria and their possible applications. FEMS
Microbiol. Rev. 22, 151–175.
De Philippis, R. and Vincenzini, M. 2003. Outermost polysac-
charidic investments of cyanobacteria: nature, significance and
possible applications. Recent Res. dev. Microbiol. 7, 13–22.
Dubois, M., Gilles, K.A., Hamilton, J.K., Rebers, P.A., and Smith,
F. 1956. Colorimetric method for determination of sugars and
related substances. Anal. Chem. 28, 350–356.
Ehling-Schulz, M., Bilger, W., and Scherer, S. 1997. UV-B-in-
duced synthesis of photoprotective pigments and extracellular
polysaccharides in the terrestrial cyanobacterium Nostoc com-
mune. J. Bacteriol. 179, 1940–1945.
Friedman, O., Dubinsky, Z., and Arad, S. 1991. Effect of light in-
tensity on growth and polysaccharides production in red and
blue-green rhodophyta unicells. Bioresour. Technol. 38, 105–110.
 Effects of light intensity on EPS components and topographical structures
Hokputsa, S., Hu, C., Paulsen, B.S., and Harding, S.E. 2003. A
physico-chemical comparative study on extracellular carbohy-
drate polymers from five desert algae. Carbohyd. Polym. 54,
27–32.
Hu, C., Liu, Y., Paulsen, B.S., Petersen, D., and Klaveness, D. 2003.
Extracellular carbohydrate polymers from five desert soil algae
with different cohesion in the stabilization of fine sand grain.
Carbohyd. Polym. 54, 33–42.
Hu, C., Liu, Y., Zhang, D., Huang, Z., and Paulsen, B.S. 2002.
Cementing mechanism of algal crusts from desert area. Chinese
Sci. Bull. 47, 1361–1368.
Huang, Z., Liu, Y., Paulsen, B.S., and Klaveness, D. 1998. Studies
on polysaccharides from three edible species of Nostoc (cyano-
bacteria) with different colony morphologies: comparison of
monosaccharide compositions and viscosities of polysaccharides
from field colonies and suspension cultures. J. Phycol. 34, 962–
968.
Li, P., Liu, Z., and Xu, R. 2001. Chemical characterisation of the
released polysaccharides from the cyanobacterium Aphanothece
halophytica GR02. J. Appl. Phycol. 13, 71–77.
Lowry, O.H., Rosebrough, N.J., Farr, A.L., and Randall, R.J. 1951.
Protein measurement with the folin phenol reagent. J. Biol.
Chem. 193, 265–275.
Mandal, S., Singh, R., and Patel, V. 2011. Isolation and character-
ization of exopolysaccharides secreted by a toxic dinoflagellate,
Amphidinium carterae Hulburt 1957 and its probable role in
harmful algal blooms (HABs). Microb. Ecol. 62, 518–527.
Moreno, J., Vargas, M.A., Olivares, H., Rivas, J., and Guerrero, M.G.
1998. Exopolysaccharides production by the cyanobacterium
Anabaena sp. ATCC 33047 in batch and continuous culture. J.
Biotechnol. 60, 175–182.
Mota, R., Guimaraes, R., Büttel, Z., Rossi, F., Colica, G., Silva, C.J.,
Santos, C., Gales, L., Zille, A., De Philippis, R., Pereira, S.B., and
et al. 2013. Production and characterization of extracellular car-
bohydrate polymer from Cyanothece sp. CCY 0110. Carbohyd.
Polym. 92, 1408–1415.
Neu, T.R. and Marshall, K.C. 1990. Bacterial polymers: physico-
chemical aspects of their interactions at interfaces. J. Biomater.
183
Appl. 5, 107–133.
Otero, A. and Vincenzini, M. 2003. Extracellular polysaccharides
synthesis by Nostoc strains as affected by N source and light
intensity. J. Biotechnol. 102, 143–152.
Pereira, S., Zille, A., Micheletti, E., Moradas-Ferreira, P., De Philippis,
R., and Tamagnini, P. 2009. Complexity of cyanobacterial exo-
polysaccharides: composition, structures, inducing factors and
putative genes involved in their biosynthesis and assembly.
FEMS Microbiol. Rev. 33, 917–941.
Pletikapic, G., Radic, T.M., Zimmermann, A.H., Svetlicic, V., Pfann-
kuchen, M., Maric, D., Godrijan, J., and Zutic, V. 2011. AFM
imaging of extracellular polymer release by marine diatom
Cylindrotheca closterium (Ehrenberg) Reiman & J.C. Lewin. J.
Mol. Recognit. 24, 436–445.
Rippka, R., Deruelles, J., Waterbury, J.B., Herdman, M., and Stanier,
R.Y. 1979. Generic assignments, strain histories and properties
of pure cultures of cyanobacteria. J. Gen. Microbiol. 111, 1–61.
Su, J., Jia, S., Chen, X., and Yu, H. 2008. Morphology, cell growth,
and polysaccharides production of Nostoc flagelliforme in liq-
uid suspension culture at different agitation rates. J. Appl.
Phycol. 20, 213–217.
Suresh Kumar, A., Mody, K., and Jha, B. 2007. Bacterial exopoly-
saccharides-a perception. J. Basic Microb. 47, 103–117.
Trabelsi, L., Ben Ouada, H., Bacha, H., and Ghoul, M. 2009.
Combined effect of temperature and light intensity on growth
and extracellular polymeric substance production by the cya-
nobacterium Arthrospira platensis. J. Appl. Phycol. 21, 405–412.
Vincenzini, M., Philippis, R., Sili, C., and Materassi, R. 1993. Stabi-
lity of molecular and rheological properties of the exopolyzsac-
charide produced by Cyanospira capsulata cultivated under dif-
ferent growth conditions. J. Appl. Phycol. 5, 539–541.
Yoshimura, H., Kotake, T., Aohara, T., Tsumuraya, Y., Ikeuchi, M.,
and Ohmori, M. 2012. The role of extracellular polysaccharides
produced by the terrestrial cyanobacterium Nostoc sp. strain
HK-01 in NaCl tolerance. J. Appl. Phycol. 24, 237–243.
Yu, H., Jia, S., and Dai, Y. 2010. Accumulation of exopolysaccha-
rides in liquid suspension culture of Nostoc flagelliforme cells.
Appl. Biochem. Biotechnol. 160, 552–560.